Fluctuating selection

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Alleles A and B fluctuate in adaptive value over time Fluctuating selection.png
Alleles A and B fluctuate in adaptive value over time

Fluctuating selection is a mode of natural selection characterized by the fluctuation of the direction of selection on a given phenotype over a relatively brief period of evolutionary time. For example, a species of plant may come in two varieties: one which prefers wetter soil and one which prefers dryer soil. During a period of wet years, the wet variety will be more fit and produce more offspring, and thereby increase the frequency of wet-preferring plants. If this wet period is followed by drought, the dry variety will be selected for and its numbers will increase. As periods of dryness and wetness fluctuate, so too does selection on dry-preferring and wet-preferring plants. Fluctuating selection is also manifest at the genic level. Consider two alleles, A and B, which are found at the same locus. Fluctuating selection dynamics are at play when selection favors A at time t0, B at t1 and A again at t2.

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Fluctuating selection has been characterized by several mathematical models. [1] [2] [3] Under some circumstances, fluctuating selection may lead to a balanced polymorphism. [4] When two species exert selection on one another, e.g. a host and its parasite, this can lead to fluctuating selection dynamics. [5] [6] [7]

Red Queen dynamics and the maintenance of sex

The Red Queen hypothesis describes coevolutionary 'arms races' between antagonistic species (predators and prey, parasites and hosts, competitors with overlapping niches), emphasizing competition between species and populations rather than within them. Under Red Queen dynamics, a species must adapt to shifting selection pressures of the ever-changing biota which constitute its environment or face extinction. Experiments in Red Queen environments on real and simulated populations have offered strong support for the maintenance of sexual reproduction despite the two-fold cost of sex. [8] [9]

Fluctuating selection may also play an important role in host-parasite coevolutionary relationships, specifically in the maintenance of sex. It has been shown that coevolutionary arms race dynamics between host and parasite give way to fluctuating selection dynamics in a minimal environment. [6] Fluctuating selection in Red Queen environments has been suggested as an explanation for the persistence of sex:

The essence of sex in our theory is that it stores genes that are currently bad but have promise for reuse. It continually tries them in combination, waiting for the time when the focus of disadvantage has moved elsewhere. When this has happened, the genotypes carrying such genes spread by successful reproduction, becoming simultaneously stores for other bad genes and thus onward in continuous succession. [8]

In this conception of sex, the population is a storehouse of variation and sex is a mechanism for distributing old, minority variants once they become useful. This theory depends on fluctuating selection, as fluctuating selection dynamics make adaptive previously maladaptive variants due to ecological shifts.

See also

Related Research Articles

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Parasitism relationship between species where one organism lives on or in another organism, causing it harm

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Ectosymbiosis Symbiosis in which the symbiont lives on the body surface of the host

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Coevolution Two or more species influencing each others evolution

In biology, coevolution occurs when two or more species reciprocally affect each other's evolution through the process of natural selection. The term sometimes is used for two traits in the same species affecting each other's evolution, as well as gene-culture coevolution.

Genetic variation The concept and mechanisms of variation in alleles of genes

Genetic variation is the difference in DNA among individuals or the differences between populations. There are multiple sources of genetic variation, including mutation and genetic recombination. Mutations are the ultimate sources of genetic variation, but other mechanisms such as sexual reproduction and genetic drift contribute to it as well.

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Evolution of sexual reproduction How sexually reproducing multicellular organisms could have evolved from a common ancestor species

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Genome size

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Red Queen hypothesis Concept in evolutionary biology

The Red Queen hypothesis, also referred to as Red Queen's, the Red Queen effect, the Red Queen model, Red Queen's race, and Red Queen dynamics, is a hypothesis in evolutionary biology which proposes that species must constantly adapt, evolve, and proliferate in order to survive while pitted against ever-evolving opposing species. The hypothesis was intended to explain the constant (age-independent) extinction probability as observed in the paleontological record caused by co-evolution between competing species; however, it has also been suggested that the Red Queen hypothesis explains the advantage of sexual reproduction at the level of individuals, and the positive correlation between speciation and extinction rates in most higher taxa.

Sex-limited genes are genes that are present in both sexes of sexually reproducing species but are expressed in only one sex and have no penetrance, or are simply 'turned off' in the other. In other words, sex-limited genes cause the two sexes to show different traits or phenotypes, despite having the same genotype. This term is restricted to autosomal traits, and should not be confused with sex-linked characteristics, which have to do with genetic differences on the sex chromosomes. Sex-limited genes are also distinguished from sex-influenced genes, where the same gene will show differential expression in each sex. Sex-influenced genes commonly show a dominant/recessive relationship, where the same gene will have a dominant effect in one sex and a recessive effect in the other. However, the resulting phenotypes caused by sex-limited genes are present in only one sex and can be seen prominently in various species that typically show high sexual dimorphism.

Host–parasite coevolution Mutually adaptive genetic change of a host and a parasite

Host–parasite coevolution is a special case of coevolution, where a host and a parasite continually adapt to each other. This can create an evolutionary arms race between them. A more benign possibility is of an evolutionary trade-off between transmission and virulence in the parasite, as if it kills its host too quickly, the parasite will not be able to reproduce either. Another theory, the Red Queen hypothesis, proposes that since both host and parasite have to keep on evolving to keep up with each other, and since sexual reproduction continually creates new combinations of genes, parasitism favours sexual reproduction in the host.

Interlocus sexual conflict is a type of sexual conflict that occurs through the interaction of a set of antagonistic alleles at two or more different loci, or the location of a gene on a chromosome, in males and females, resulting in the deviation of either or both sexes from the fitness optima for the traits. A co-evolutionary arms race is established between the sexes in which either sex evolves a set of antagonistic adaptations that is detrimental to the fitness of the other sex. The potential for reproductive success in one organism is strengthened while the fitness of the opposite sex is weakened. Interlocus sexual conflict can arise due to aspects of male–female interactions such as mating frequency, fertilization, relative parental effort, female remating behavior, and female reproductive rate.

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<i>Hamiltosporidium</i>

Hamiltosporidium is a genus of Microsporidia, which are intracellular and unicellular parasites. The genus, proposed by Haag et al. in 2010, contains two species; Hamiltosporidium tvaerminnensis, and Hamiltosporidium magnivora. Both species infect only the crustacean Daphnia magna (Waterflea).

Josephine Pemberton British evolutionary biologist

Josephine M. Pemberton is a British evolutionary biologist. She is Chair of Natural History at the University of Edinburgh, where she conducts research in parentage analysis, pedigree reconstruction, inbreeding depression, parasite resistance, and quantitative trait locus (QTL) detection in natural populations. She has worked primarily on long-term studies of soay sheep on St Kilda, and red deer on the island of Rùm.

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